FEEDING HABITS OF THE GENET GENETTA GENETTA IN AN IBERIAN CONTINENTAL WETLAND M

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FEEDING HABITS OF THE GENET GENETTA GENETTA IN AN IBERIAN CONTINENTAL WETLAND

M

AITE

SÁNCHEZ

¹

, P

ATRICIA

RODRIGUES

¹

, V

ICENTE

ORTUÑO

²

, J

UAN

HERRERO

^3*

1Department of Ecology. University of Alcalá. E-28871 Alcalá de Henares, Spain e-mail: maite.soy@gmail.com; patriciarodrigues@ua.pt

2Department of Animal Biology and Physical Anthropology. University of Alcalá, E-28871 Alcalá de Henares, Spain; e-mail: vicente.ortuno@uah.es

3Department of Ecology. University of Alcalá. E-28871 Alcalá de Henares. Ega Wildlife Consultants. Sierra de Vicort 31. E-50003 Zaragoza, Spain

*Corresponding author: juan.herrero@uah.es, egasl@arrakis.es

Received 19 August 2008; accepted 24 November 2008

ABSTRACT - This study quantified the diet of the genet Genetta genetta in an Iberian con- tinental wetland, the Galachos Nature Reserve, northern Spain, based on the frequency with which items appeared in five monthly-surveyed latrines during 2004-2005. Prey types were identified de visu using identification keys and expert advice. The genet was confirmed as an opportunistic and generalist predator, its diet including mammals (95.0%), plants (68.3%), and arthropods (60.0%) as main prey. With the exception of a newly available prey species, the alien crayfish Procambarus clarkii, the genet probably consumed arthro- pods because of chance encounters rather than active search. The consumption of fruits and small mammals varied seasonally.

Key words: Seasonality, Apodemus sylvaticus, Mus spretus, protected areas, Procambarus clarkii, Spain

RIASSUNTO - Alimentazione della genetta Genetta genetta in un’area umida continen- tale della penisola iberica. La dieta della genetta è stata investigata in un’area umida inter- na della Spagna settentrionale, la Riserva Naturale dei Galachos. La frequenza di ciascuna categoria alimentare è stata espressa come numero di occorrenze mensili in cinque latrine monitorate per il periodo 2004-2005. Le categorie sono state identificate de visu tramite chiavi per il riconoscimento o l’intervento di singoli esperti. I risultati confermano il com- portamento alimentare generalista e opportunista della genetta. La dieta ha incluso princi- palmente mammiferi (95%), vegetali (68,3%) e artropodi (60%). Questi ultimi, con l’eccezione del gambero Procambarus clarkii, introdotto e di recente presenza, sono proba- bilmente utilizzati in modo opportunistico, piuttosto che in seguito a ricerca attiva. L’uso dei frutti e dei micromammiferi varia stagionalmente.

Parole chiave: Variazioni stagionali, Apodemus sylvaticus, Mus spretus, aree protette, Pro- cambarus clarkii, Spagna

INTRODUCTION

The genet Genetta genetta is a me- dium-sized, carnivorous viverrid which

is native to northern Africa. It was in- troduced into southern Europe several centuries ago and today its populations occupy a wide area on both continents

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(Delibes, 1999). The IUCN (2007) con- sider the species to be of “Least Con- cern”.

The diet is the aspect of genet biology which has been most investigated (Li- vet and Roeder, 1987; Hamdine et al., 1993; Ruiz-Olmo and López-Martín, 1993; Ballesteros et al., 2003), in part because faeces can be easily gathered from latrines. Studies on genet diet have covered a wide geographical range, including northern Africa (De- libes et al., 1989; Hamdine et al., 1993), France (Chanudet et al., 1967;

Cugnasse and Riols, 1984; Livet and Roeder, 1987; Lodé et al., 1991; Le Jaques and Lodé, 1994), Portugal (Rosalino and Santos-Reis, 2002), and Spain (Delibes, 1974; Calviño et al., 1984; 1987; Ruiz-Olmo and López- Martín, 1993, 1996; Virgós et al., 1999; Arrizabalaga et al., 2002; Balles- teros et al., 2003; Torre et al., 2003).

These studies have shown that the genet is a generalist and opportunistic predator (Delibes et al., 1989; Calviño et al., 1984; Ruiz-Olmo and López- Martín, 1993; Le Jaques and Lodé, 1994; Rosalino and Santos-Reis, 2002).

Small mammals, particularly the wood mouse Apodemus sylvaticus, are the most important component of the diet (Delibes, 1974; Delibes et al., 1989;

Palomares and Delibes, 1991; Hamdine et al., 1993; Le Jaques and Lodé, 1994;

Lodé et al., 1991; Ballesteros et al., 2003).

Ruíz-Olmo (1993) classified the species as a small-mammal specialist predator, even though its feeding behaviour can vary according to local and seasonal changes in the availability of prey types (Virgós et al., 1999).

The diet of the genet in wetlands has been

the object of a few studies (Ruiz-Olmo and López-Martín, 1993; Rosalino and Santos-Reis, 2002). In our study, we analysed the diet of the genet in the middle River Ebro Valley, northern Iberia, with the aim of identifying the main prey types and their seasonal variation.

STUDY AREA

The study area covered 751 ha within the Galachos Nature Reserve (GNR), a wetland in the Middle Ebro Valley, near Saragossa, Aragon, Spain (41° 38' 46’’, 08 North, 0°

53' 33’’, 51 West). The reserve mainly con- sists of riparian forests (42%) and agricul- tural lands (27%) (Herrero et al., 2006). In 1994, the GNR was designated as a Special Protection Area under the European Bird Directive because of the high diversity of birds that breed there, particularly night heron Nycticorax nycticorax. In 1998, it was designated a Site of Community Im- portance and included in the Natura 2000 Network. The climate is semi-arid Mediter- ranean (Rivas and Baselga, 2005), annual rainfall averaging 272 mm.

In the GNR, carnivores include red fox Vulpes vulpes, badger Meles meles, stone marten Martes foina, weasel Mustela nivalis and otter Lutra lutra. Other me- dium-sized mammals are European rabbit Oryctolagus cuniculus and Iberian hare Lepus granatensis.

METHODS

In the first week of each month from July 2004 to June 2005, genet scats were collected from five latrines. The latrine/month was considered as the sample unit. Each month, on average, two fresh scats were removed from each latrine, stored in plastic bags, and kept frozen until analyses. Each sample was thawed and added to water with soap and alcohol to allow its disinte-

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gration and avoid the breaking of diagnostic elements. Once the samples were thor- oughly soaked, their constituents were separated by pincers in a dry tray.

Hemimandibles and hemimaxilles were used to identify small mammals eaten by the genet (Gállego and Alemany, 1985;

Gállego and López, 1985), and were also compared with a reference collection.

Birds’ feathers were identified by two groups of ornithologists, to allow the com- parison of their results. Plant matter was identified using botanical keys (Aizpuru et al., 1999) and the floristic catalogue of the GNR (Regato, 1988). Arthropods were identified de visu (crayfish) or using avail- able keys (Paulian and Baraud, 1982; Bar- rientos and Ferrández, 1985; Martínez et al., 1985; Chinery, 1997; Baraud, 1992;

Vives, 2000). For each item the frequency of occurrence (F%) was calculated as the number of latrines containing the item di- vided by the total number of latrines/month x 100. Microlepidoptera and dermestid larvae (Coleoptera), also ants (Tapinoma, Formicidae, Hymenoptera), were considered contamination of the excrements of the latrine and thus excluded from the analysis.

Seasonal (winter: XII-II, spring: III-V, summer: VI-VIII, autumn: IX-XI) changes in the frequency of use of the main prey types (plant matter, fruits and seeds, crustaceans, arthropods, birds and mammals) were evaluated using the Ȥ² test. The p–

value for statistical significance was calculated using Holm’s sequential Bonferroni method, accepting an overall ʌ value of 0.05 (Rice, 1989). Small mammal consumption, determined through the count of hemimandibles and hemimaxilles, was de- scribed on a monthly basis and analysed seasonally.

RESULTS

We collected a total of 135 faecal samples, containing at least 36 different food items (Tab. 1).

The bulk of genet diet consisted of mammals (F% = 95%), particularly wood mouse Apodemus sylvaticus (55.0%), Western Mediterranean mouse Mus spretus (40%), and greater white-toothed shrew Crocidura russula (26.7%). Other mammals included black rat Rattus rattus (5.0%) and voles (Microtus sp.; 3.3%).

Plant matter, particularly Poaceae (26.7%), was common (68.3%), and, in many cases, appeared as an undigested distal branch of faeces. Arthropods, which occurred in 60.0% of the samples, included mainly crayfish (43.3%) and insects (46.7%). The invertebrates most preyed upon included Coleoptera (28.3%) - Dorcus parallelepipedus (10.0%) and Amphimalon sp. (10.0%) -, Orthoptera (21.7%), and large spiders (21.7%). Fruits (58.3%) mostly con- sisted of figs Ficus carica (18.3%), whilst Passeriformes (12 species) - es- pecially blackbird Turdus merula (6.7%) -, prevailed among birds (46.7%). The frequency of use of fruits and seeds varied significantly among seasons (Ȥ² = 13.0, d.f.=3, P < 0.01), showing a peak in autumn (Tab. 2).

The consumption of small mammals varied seasonally (Ȥ² = 226, d.f.=3, P <

0.001), rodents being less frequent in genet diet from April through Novem- ber (Fig. 1).

DISCUSSION

In the GNR, the genet was confirmed as a generalist and opportunistic predator. Its main prey were rodents living in densely vegetated habitats, and crayfish. This preference, together with the frequent presence in the scats of the female flowers of willows and poplar cat-

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Table 1 - Diet of the genet (F%) in the Galachos Nature Reserve, northern Spain, based on monthly faecal samples collected from five latrines in 2004-2005 (N = 60 latrines).

FOOD ITEMS F% N

PLANT MATTER 68.3 41

Leaves 50.0 30

Compositae 3.3 2

Poaceae 26.7 16

Rosaceae 1.7 1

Salicaceae 1.7 10

Salix sp. 10.0 6

Tamarix gallica 16.7 10

Poligonum sp. 1.7 1

Fruits and seeds 58.3 35

Salicaceae 1.7 1

Populus sp. (cataphylls) 1.7 1

Salix sp. 5.0 3

Moraceae 18.3 11

Ficus carica 18.3 11

Poaceae 5.0 3

Zea mays 1.7 1

Triticum durum 3.3 2

Polygonaceae 16.7 10

Polygonum sp. 10.0 6

Undetermined fruits and seeds 5.0 3

ANIMAL MATTER 98.3 59

Invertebrates 61.7 37

Arthropoda 60.0 36

Aracnida 21.7 13

Procambarus clarkii 43.3 26

Insects 46.7 28

Coleoptera 28.3 17

Cerambycidae 3.3 2

Lamia textor 1.7 1

Ergates faber 1.7 1

Scarabaeidae (Dynastinae) 3.3 2

Phyllognathus excavatus 1.7 1

Oryctes nasicornis 1.7 1

Scarabeidae (Melolonthinae) 11.7 7

Amphimallon sp. 10.0 6

Lucanidae Dorcus parallelipipedus 10.0 6 Dermaptera Forcipula sp. 3.3 2

Orthoptera 21.7 13

Acrididae 21.7 13

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Table 1 - continues

FOOD ITEMS F% N

Gryllotalpa gryllotalpa 5.0 3

Undetermined insects 1.7 1

Lepidoptera larvae 13.3 8

Undetermined larvae 1.7 1

Gasteropoda 1.7 1

Vertebrates 96.7 58

Birds 46.7 28

Passeriformes 25.0 15

Aegithalos caudatus 1.7 1

Dendrocopos major 1.7 1

Erithacus rubecula 1.7 1

Luscinia megarhynchos 1.7 1

Passer sp. 1.7 1

Phylloscopus collybita 1.7 1

Sturnus sp. 3.3 2

Sturnus unicolor 3.3 2

Sylvia atricapilla 1.7 1

Turdus merula 6.7 4

Turdus philomelos 1.7 1

Vanellus vanellus 1.2 1

Undetermined birds 21.7 13

Mammals 95.0 57

Crocidura russula 26.7 16

Rodentia 56.7 34

Apodemus sylvaticus 55.0 33

Microtus sp. 3.3 2

Mus spretus 40.0 24

Rattus rattus 5.0 3

Rattus sp. 8.3 5

Undetermined mammals 1.7 1

kins, suggests that riparian forests may be the genet‘s preferred habitat for seeking food.

Whilst in wetlands of central Portugal (Rosalino and Santos-Reis, 2002) crayfish were very rare in genet diet (<1%

of 150 faecal samples), in the GNR crayfish are frequently preyed upon also by wild boar Sus scrofa (Herrero et al., 2006), confirming the importance of this alien species as prey for multiple

secondary consumers in the Iberian Peninsula (see also Pedroso and San- tos-Reis, 2007 about the otter).

Among arthropods, insects often are the second most important prey in genet diet (Delibes et al., 1989; Ham- dine et al., 1993; Rosalino and Santos, 2002). In the GNR insects were mostly large, above-ground dwelling, crepus- cular species, including many orb web spiders, grasshoppers and beetles, closely

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Table 2 - Seasonal variation (F%) of the main items in the diet of the genet in the Galachos Nature Reserve; N. latrines/season: 15; *significant.

Food items Winter Spring Summer Autumn Ȥ² P Fruits and seeds 20.0 13.3 33.3 86.7 13 0.005*

Arthropoda 26.7 26.7 66.7 13.3 7.2 0.066

Crustaceans 66.7 40.0 26.7 20.0 5 0.17

Plant matter 46.7 60.0 80.0 53.3 1.5 0.67

Birds 66.7 60.0 53.3 40.0 1.06 0.79

Mammals 100.0 86.7 53.3 93.3 1.06 0.79

0 20 40 60 80 100

J F M A M J J A S O N D

N

Figure 1 - Number (N) of hemimandibles and hemimaxillas of small mammals found in monthly faecal samples collected from 5 latrines in 2004-2005 (N = 60 latrines).

associated with the vegetation of the area. We suggest that genets consumed them by chance, and not through active searching. The absence of cryptic larvae (living in roots or wood), such as those of some beetles, supports this interpretation.

In some genet populations, birds, mainly thrushes (Turdidae) and other passerines, can represent an important food resource (Le Jacques and Lodé,

1994; Cugnasse and Riols, 1984). In our study area, the genet consumed a wide variety of birds. The presence of a lapwing Vanellus vanellus well out of its migratory period suggests that genets rely on defenceless individuals, either young or weakened.

Plant matter was a frequent item in the diet of the genet, but apart from figs, which can represent an important re- source (Ballesteros et al., 2003), it

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mainly consisted of leaves, particularly those of Poaceae. These appeared un- digested, with unaltered structure and colour, suggesting they were not consumed for their nutritional value. As with other carnivores, genets might eat Poaceae to aid digestion, help to elimi- nate hair from the intestine, induce vomiting to excrete ingested toxins, alleviate throat and stomach inflamma- tions, or as a source of folic acid (Mor- ris, 1996).

Genet diet varies seasonally (Delibes, 1974; Cugnasse and Riols, 1984;

Calviño et al., 1984; Delibes et al., 1989; Palomares and Delibes, 1991;

Lodé et al., 1991; Hamdine et al., 1993; Ruiz-Olmo and López-Martín, 1993; Le Jaques and Lodé, 1994; Rosa- lino and Santos-Reis, 2002; Ballesteros et al., 2003), according to the variation of the availability of its various prey or other environmental factors, such as rainfall (Le Jaques and Lodé, 1994). As with previous studies, in the GNR fruit consumption peaked in autumn (Calviño et al., 1984; Cugnasse and Riols, 1984; Lodé et al., 1991), whilst small mammals were the most common prey in winter (Calviño et al., 1984;

Rosalino and Santos-Reis, 2002; Ball- esteros et al., 2003), probably because of their relatively higher availability compared to fruits and Arthropoda in the cold season. The other items did not show any significant variation, particularly birds, which many studies (Calviño et al., 1984; Cugnasse and Riols, 1984; Lodé et al., 1991; Le Jaques and Lodé, 1994) have reported to be more frequent in winter and spring, coinciding with the breeding seasons of many species at our lati- tudes.

The pattern of seasonal variation found in the GNR with respect to other studies could be due either to environmental factors, such as climate and prey availability, or to sample size.

In general, our results are similar to those of previous studies based on the analysis of gastric contents or faeces (Braña and Del Campo, 1982; Delibes et al. 1989; Hamdine et al., 1993;

Rosalino and Santos, 2002; Le Jacques and Lodé, 1994), even though they sometimes sharply differ in their methods and sample size (36 in Braña and Del Campo, 1982; 150 in Torre et al., 2003; more than 2000 in Lodé et al., 1991), making comparisons rather dif- ficult. For instance, the extraction of diagnostic elements from faeces can be achieved using re-hydrated, oven-dried excrements (Palomares and Delibes, 1991; Lodé et al. 1991), faecal samples hydrated with different dilutions (De- libes et al., 1989; Ruíz-Olmo and López Martín, 1993, 1996; Arriza- balaga et al., 2002; Roberts et al. 2007) or with no previous treatment (Calviño et al., 1984; Ballesteros et al., 2003). In some cases, the method used is either partially (Hamdine et al., 1993) or not at all explained (Rosalino and Santos- Reis, 2002; Torre et al., 2003). Finally, the deterioration of the diagnostic elements is a main problem of faecal analyses. Further research on genet feeding behaviour in wetlands is needed to better understand the species on which the genet preys.

ACKNOWLEDGEMENTS

We thank the ranger F. Sebastián who collected the faeces. Ch. González, J.

M. Sánchez (La Alfranca Fauna Re-

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covery Centre, Government of Aragon) and J. Blasco, (bird-ringer) provided de visu identification of feathers. A. Al- dezabal (Basque Country University) provided de visu plant identification.

Philippe Gaubert (Systematics and Evolution Department of the French Natural History Museum) provided bibliography. Carlos Nores (Oviedo University) and Juan Carlos Blanco (wildlife consultant) reviewed an early draft of the manuscript.

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